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de Waal, F. B., & Seres, M. (1997). Propagation of handclasp grooming among captive chimpanzees. Am. J. Primatol., 43(4), 339–346.
Abstract: A grooming posture previously reported for two wild chimpanzee (Pan troglodytes) communities developed spontaneously in a captive group of the same species. This offered a unique opportunity to follow the propagation of a new social custom. The posture consists of two partners grasping hands--either both right hands or both left hands--and raising the arms in an A-frame above their heads while mutually grooming with their free hands. The propagation of this pattern was followed over a 5 year period. In the beginning, handclasps were always initiated by the same adult female. This female initiated the posture mainly with her adult female kin. In subsequent years, these relatives became frequent participants in the posture with each other as well as with nonrelatives. Over the years the posture increased in frequency and duration and spread to the majority of adults and also to a few adolescents and older juveniles. The pattern persisted after removal of the apparent originator.
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Aureli, F., & de Waal, F. B. (1997). Inhibition of social behavior in chimpanzees under high-density conditions. Am. J. Primatol., 41(3), 213–228.
Abstract: This is the first study to investigate the short-term effects of high population density on captive chimpanzees (Pan troglodytes). Subjects of the study were 45 chimpanzees living in five different groups at the Yerkes Regional Primate Research Center. The groups were observed under two conditions: 1) when they had access to both the indoor and outdoor sections of their enclosures; 2) during cold days when they were locked into the indoor runs, which reduced the available space by more than half. Under the high-density condition, allogrooming and submissive greetings decreased, but juvenile play increased. Remarkably, the rate of various forms of agonistic behavior, such as aggression, bluff charge, bluff display, and hooting, occurred less frequently under the high-density condition. This general decrease in adult social activity, including agonistic behavior, can be interpreted as an inhibition strategy to reduce opportunities for conflict when interindividual distances are reduced. This strategy is probably effective only in the short run, however. Behavioral indicators of anxiety, such as rough scratching and yawning, showed elevated rates, suggesting increased social tension under the high-density condition.
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Koski, S. E., Koops, K., & Sterck, E. H. M. (2007). Reconciliation, relationship quality, and postconflict anxiety: testing the integrated hypothesis in captive chimpanzees. Am. J. Primatol., 69(2), 158–172.
Abstract: Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The “integrated hypothesis” links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.
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Langergraber, K., Mitani, J., & Vigilant, L. (2009). Kinship and social bonds in female chimpanzees (Pan troglodytes). Am. J. Primatol., 71(10), 840–851.
Abstract: A large body of theoretical and empirical research suggests that kinship influences the development and maintenance of social bonds among group-living female mammals, and that human females may be unusual in the extent to which individuals form differentiated social relationships with nonrelatives. Here we combine behavioral observations of party association, spatial proximity, grooming, and space use with extensive molecular genetic analyses to determine whether female chimpanzees form strong social bonds with unrelated individuals of the same sex. We compare our results with those obtained from male chimpanzees who live in the same community and have been shown to form strong social bonds with each other. We demonstrate that party association is as good a predictor of spatial proximity and grooming in females as it is in males, that the highest party association indices are consistently found between female dyads, that the sexes do not differ in the long-term stability of their party association patterns, and that these results cannot be explained as a by-product of the tendency of females to selectively range in particular areas of the territory. We also show that close kin (i.e. mother-daughter and sibling dyads) are very rare, indicating that the vast majority of female dyads that form strong social bonds are not closely related. Additional analyses reveal that “subgroups” of females, consisting of individuals who frequently associate with one another in similar areas of the territory, do not consist of relatives. This suggests that a passive form of kin-biased dispersal, involving the differential migration of females from neighboring communities into subgroups, was also unlikely to be occurring. These results show that, as in males, kinship plays a limited role in structuring the intrasexual social relationships of female chimpanzees.
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Shoshani, J., Kupsky, W. J., & Marchant, G. H. (2006). Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull, 70(2), 124–157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
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Dunbar, R. I. M., McAdam, M. R., & O'connell, S. (2005). Mental rehearsal in great apes (Pan troglodytes and Pongo pygmaeus) and children. Behav. Process., 69(3), 323–330.
Abstract: The ability to rehearse possible future courses of action in the mind is an important feature of advanced social cognition in humans, and the “social brain” hypothesis implies that it might also be a feature of primate social cognition. We tested two chimpanzees, six orangutans and 63 children aged 3-7 years on a set of four puzzle boxes, half of which were presented with an opportunity to observe the box before being allowed to open it (“prior view”), the others being given without an opportunity to examine the boxes before handling them (“no prior view”). When learning effects are partialled out, puzzle boxes in the “prior view” condition were opened significantly faster than boxes given in the “no prior view” condition by the children, but not by either of the great apes. The three species differ significantly in the speed with which they opened boxes in the “no prior view” condition. The three species' performance on this task was a function of relative frontal lobe volume, suggesting that it may be possible to identify quantitative neuropsychological differences between species.
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Hirata, S. (2007). A note on the responses of chimpanzees (Pan troglodytes) to live self-images on television monitors. Behav. Process., 75(1), 85–90.
Abstract: The majority of studies on self-recognition in animals have been conducted using a mirror as the test device; little is known, however, about the responses of non-human primates toward their own images in media other than mirrors. This study provides preliminary data on the reactions of 10 chimpanzees to live self-images projected on two television monitors, each connected to a different video camera. Chimpanzees could see live images of their own faces, which were approximately life-sized, on one monitor. On the other monitor, they could see live images of their whole body, which were approximately one-fifth life-size, viewed diagonally from behind. In addition, several objects were introduced into the test situation. Out of 10 chimpanzees tested, 2 individuals performed self-exploratory behaviors while watching their own images on the monitors. One of these two chimpanzees successively picked up two of the provided objects in front of a monitor, and watched the images of these objects on the monitor. The results indicate that these chimpanzees were able to immediately recognize live images of themselves or objects on the monitors, even though several features of these images differed from those of their previous experience with mirrors.
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Vokey, J. R., Rendall, D., Tangen, J. M., Parr, L. A., & de Waal, F. B. M. (2004). Visual kin recognition and family resemblance in chimpanzees (Pan troglodytes). J Comp Psychol, 118(2), 194–199.
Abstract: The male-offspring biased visual kin recognition in chimpanzees (Pan troglodytes) reported by L. A. Parr and F. B. M. de Waal (1999) was replicated with human (Homo sapiens) participants and a principal components analysis (PCA) of pixel maps of the chimpanzee face photos. With the same original materials and methods, both humans and the PCA produced the same asymmetry in kin recognition as found with the chimpanzees. The PCA suggested that the asymmetry was a function of differences in the distribution of global characteristics associated with the framing of the faces in the son and daughter test sets. Eliminating potential framing biases, either by cropping the photos tightly to the faces or by rebalancing the recognition foils, eliminated the asymmetry but not human participants' ability to recognize chimpanzee kin.
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Flack, J. C., Jeannotte, L. A., & de Waal, F. B. M. (2004). Play signaling and the perception of social rules by juvenile chimpanzees (Pan troglodytes). J Comp Psychol, 118(2), 149–159.
Abstract: Prescriptive social rules are enforced statistical regularities. The authors investigated whether juvenile chimpanzees (Pan troglodytes) recognize and use enforced statistical regularities to guide dyadic play behavior. They hypothesized (a) that proximity of adults, especially mothers of younger play partners, to play bouts will increase the play signaling of older partners and (b) that when juvenile-juvenile play bouts occur in proximity to adults, older partners will play at a lower intensity than when no adults are present. They found that older and younger partners increase their play signaling in the presence of the mothers of younger partners, particularly as the intensity of play bouts increases. In contrast to their hypothesis, older partners played more roughly when the mothers of younger partners were in proximity. These results suggest that juvenile chimpanzees increase play signaling to prevent termination of the play bouts by mothers of younger partners.
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Weaver, A., & de Waal, F. B. M. (2002). An index of relationship quality based on attachment theory. J Comp Psychol, 116(1), 93–106.
Abstract: Two measures are reported of the nature or quality of a mother-offspring (MO) relationship during development using brown capuchin monkeys (Cebus apella) as models. One is a qualitative classification of MO relationships as secure, resistant, or avoidant attachments. The other is an empirical ratio of relative affiliation to agonism called the MO relationship quality, or MORQ, Index. The two methods tapped similar relationship features so relationships high or low of a median split of MORQ values were heuristically labeled secure (n = 22) or insecure (n = 16), respectively. A comparison revealed extensive behavioral differences between secure and insecure MO relationships and suggested MORQ provided an objective, continuous measure of attachment security.
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